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<document id="7902F91B6700AB39431B347B254CA30E" ID-CLB-Dataset="272041" ID-DOI="10.1093/zoolinnean/zlad013" ID-GBIF-Dataset="effa6181-8326-4c7d-8c91-809f7fabb862" ID-ISSN="0024-4082" ID-Zenodo-Dep="8432389" ID-ZooBank="C3E2DFF9-A0A6-41EA-A149-0F73A2BEE5E6" IM.bibliography_approvedBy="valdenar" IM.illustrations_approvedBy="valdenar" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_approvedBy="valdenar" IM.taxonomicNames_approvedBy="valdenar" IM.treatments_approvedBy="valdenar" checkinTime="1696939870042" checkinUser="plazi" docAuthor="Silva, Felipe De Vasconcelos, Pola, Marta &amp; Cervera, Juan Lucas" docDate="2023" docId="03A6878CFF9B6519FFECFBA7EB6AF9EB" docLanguage="en" docName="zlad013.pdf" docOrigin="Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 199 (2)" docSource="http://dx.doi.org/10.1093/zoolinnean/zlad013" docStyle="DocumentStyle:4F230B9370E98E256D973D6DFB57F36C.1:ZoolJLinnSoc.2023-.journal_article" docStyleId="4F230B9370E98E256D973D6DFB57F36C" docStyleName="ZoolJLinnSoc.2023-.journal_article" docStyleVersion="1" docTitle="Tritoniidae Lamarck 1809" docType="treatment" docVersion="3" lastPageNumber="459" masterDocId="FF9FFFF4FF906517FC52FF84EE67FF8B" masterDocTitle="A stomach plate to divide them all: a phylogenetic reassessment of the family Tritoniidae (Nudibranchia: Cladobranchia)" masterLastPageNumber="476" masterPageNumber="445" pageNumber="456" updateTime="1697044790341" updateUser="ExternalLinkService">
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<mods:title id="E8DA9C59AE82223ECC9A3FC552F3F61A">A stomach plate to divide them all: a phylogenetic reassessment of the family Tritoniidae (Nudibranchia: Cladobranchia)</mods:title>
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<mods:namePart id="24EB2D98A11736D38A8464F5A80D2D86">Silva, Felipe De Vasconcelos</mods:namePart>
<mods:affiliation id="E51FD10DFD10FD0ECF835490B70E45CB">Departamento de Biología, Facultad de Ciencias del Mar y Ambientales; Universidad de Cádiz; Campus de Excelencia Internacional del Mar (CEI · MAR), Av. República Saharaui, s / n, Ap. 40. 11510 Puerto Real (Cádiz), Spain</mods:affiliation>
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<mods:namePart id="8C8AD24E8E3A2B15DAA25D90B0917B0D">Pola, Marta</mods:namePart>
<mods:affiliation id="EB2704759725ECEE0C5E3FBC88B387E9">Departamento de Biología, Facultad de Ciencias, Universidad Autónoma de Madrid; Campus de Excelencia Internacional UAM + CSIC, C / Darwin, 2, 28049 Madrid, Spain</mods:affiliation>
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<mods:namePart id="A5BD8D1610BFA310FD40B9AEA93C1AE1">Cervera, Juan Lucas</mods:namePart>
<mods:affiliation id="095CA2C3530BC98E9E388D32E0C2B21D">Departamento de Biología, Facultad de Ciencias del Mar y Ambientales; Universidad de Cádiz; Campus de Excelencia Internacional del Mar (CEI · MAR), Av. República Saharaui, s / n, Ap. 40. 11510 Puerto Real (Cádiz), Spain &amp; Instituto Universitario de Investigación Marina (INMAR), Universidad de Cádiz; Campus de Excelencia Internacional del Mar (CEI · MAR), Universidad de Cádiz, Av. República Saharaui, s / n, Ap. 40. 11510 Puerto Real (Cádiz), Spain</mods:affiliation>
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<treatment id="03A6878CFF9B6519FFECFBA7EB6AF9EB" ID-DOI="http://doi.org/10.5281/zenodo.8432393" ID-Zenodo-Dep="8432393" LSID="urn:lsid:plazi:treatment:03A6878CFF9B6519FFECFBA7EB6AF9EB" httpUri="http://treatment.plazi.org/id/03A6878CFF9B6519FFECFBA7EB6AF9EB" lastPageId="14" lastPageNumber="459" pageId="11" pageNumber="456">
<subSubSection id="C3156511FF9B651CFFECFBA7ED3FFB02" pageId="11" pageNumber="456" type="nomenclature">
<paragraph id="8BB0369AFF9B651CFFECFBA7EB78FBB6" blockId="11.[958,1311,1059,1085]" box="[958,1311,1059,1085]" pageId="11" pageNumber="456">
<heading id="D0F881F6FF9B651CFFECFBA7EB78FBB6" bold="true" box="[958,1311,1059,1085]" centered="true" fontSize="11" level="2" pageId="11" pageNumber="456" reason="3">
<emphasis id="B97BEA88FF9B651CFFECFBA7EB78FBB6" bold="true" box="[958,1311,1059,1085]" pageId="11" pageNumber="456">
Family
<taxonomicName id="4C0F4D19FF9B651CF858FBA7EB78FBB6" ID-CoL="7NLFJ" authority="Lamarck 1809" authorityName="Lamarck" authorityYear="1809" box="[1034,1311,1059,1085]" class="Gastropoda" family="Tritoniidae" kingdom="Animalia" order="Nudibranchia" pageId="11" pageNumber="456" phylum="Mollusca" rank="family">Tritoniidae Lamarck 1809</taxonomicName>
</emphasis>
</heading>
</paragraph>
<paragraph id="8BB0369AFF9B651CFF78FBD6ED3FFB02" blockId="11.[810,1459,1106,1255]" pageId="11" pageNumber="456">
<emphasis id="B97BEA88FF9B651CFF78FBD6EDCAFBE1" box="[810,941,1106,1130]" italics="true" pageId="11" pageNumber="456">Duvaucelidae</emphasis>
Iredale and ODonoghue 1923, Bull. Zool., 20: 272.
</paragraph>
</subSubSection>
<subSubSection id="C3156511FF9B651CFF78FB14EB25FB6C" pageId="11" pageNumber="456" type="reference_group">
<paragraph id="8BB0369AFF9B651CFF78FB14EDF2FB43" blockId="11.[810,1459,1106,1255]" pageId="11" pageNumber="456">
<taxonomicName id="4C0F4D19FF9B651CFF78FB14EA54FB23" authority="Odhner 1936" authorityName="Odhner" authorityYear="1936" box="[810,1075,1168,1192]" class="Gastropoda" family="Aranucidae" higherTaxonomySource="CoL,GBIF" kingdom="Animalia" order="Nudibranchia" pageId="11" pageNumber="456" phylum="Mollusca" rank="family">
<emphasis id="B97BEA88FF9B651CFF78FB14EDFDFB23" box="[810,922,1168,1192]" italics="true" pageId="11" pageNumber="456">Aranucidae</emphasis>
<bibRefCitation id="EF9E4B6BFF9B651CFFF0FB14EA54FB23" author="Odhner NH" box="[930,1075,1168,1192]" pageId="11" pageNumber="456" refId="ref26584" refString="Odhner NH. Nudibranchia dendronotacea: a revision of the system. Brussels: Musee Royal d'Histoire Naturelle de Belgique, 1936." type="book" year="1936">Odhner 1936</bibRefCitation>
</taxonomicName>
, Mém. Mus. R. his. Nat. Belg, 2(3): 1090, pl.1.
</paragraph>
<paragraph id="8BB0369AFF9B651CFF78FB4BEB25FB6C" blockId="11.[810,1459,1106,1255]" box="[810,1346,1231,1255]" pageId="11" pageNumber="456">
<taxonomicName id="4C0F4D19FF9B651CFF78FB4BEA25FB6C" authority="Odhner 1968" authorityName="Odhner" authorityYear="1968" box="[810,1090,1231,1255]" class="Gastropoda" family="Marianinidae" higherTaxonomySource="CoL,GBIF" kingdom="Animalia" order="Nudibranchia" pageId="11" pageNumber="456" phylum="Mollusca" rank="family">
<emphasis id="B97BEA88FF9B651CFF78FB4BEDD7FB6C" box="[810,944,1231,1255]" italics="true" pageId="11" pageNumber="456">Marianinidae</emphasis>
Odhner 1968
</taxonomicName>
, Ark. Zool., 20(13): 253.
</paragraph>
</subSubSection>
<subSubSection id="C3156511FF9B6519FF78FA8DEB6AF9EB" lastPageId="14" lastPageNumber="459" pageId="11" pageNumber="456" type="diagnosis">
<paragraph id="8BB0369AFF9B651CFF78FA8DEBEDFAD4" blockId="11.[810,1459,1288,1375]" pageId="11" pageNumber="456">
<emphasis id="B97BEA88FF9B651CFF78FA8DEDF7FAAB" box="[810,912,1289,1312]" italics="true" pageId="11" pageNumber="456">Diagnosis:</emphasis>
Rhinophore palmate retractable and sheathed, club smooth surrounded by a distal circle of vertically branched processes. One row of dorsolateral, arborescent secondary gills.
</paragraph>
<paragraph id="8BB0369AFF9B651CFF78FA04EA74F89B" blockId="11.[810,1460,1408,1808]" pageId="11" pageNumber="456">
<emphasis id="B97BEA88FF9B651CFF78FA04EDCEFA13" box="[810,937,1408,1432]" italics="true" pageId="11" pageNumber="456">Morphology:</emphasis>
Body elongated and slightly quadrangular. Notum smooth or tuberculate. Oral veil entire or bilobed, with simple or branched digitiform velar processes. Outermost processes are ventrally grooved tentacles. Rhinophore sheath tubular, with margins smooth, undulated or with one or more processes. Rhinophore retractile and palmate formed by a smooth club surrounded by a distal circle of vertical branched processes. Secondary gills project dorsolateral. Radular formula n.1.1.1.n or n.1.n. Rachidian tooth uni- or tricuspid. Masticatory borders of the jaws denticulate, differentiated or smooth. True stomach plates absent or present. ODG complex holohepatic, with right and leħ lobe fused, or cladohepatic, with right and leħ lobe separated. Penis unarmed.
</paragraph>
<paragraph id="8BB0369AFF9B6519FF78F8B6ECFAFA68" blockId="11.[810,1459,1842,1991]" lastBlockId="14.[129,778,1451,1977]" lastPageId="14" lastPageNumber="459" pageId="11" pageNumber="456">
<emphasis id="B97BEA88FF9B651CFF78F8B6EDEFF8C1" box="[810,904,1842,1866]" italics="true" pageId="11" pageNumber="456">Remarks:</emphasis>
The traditional family
<taxonomicName id="4C0F4D19FF9B651CF8CAF8B6ED0EF8E2" authority="(sensu Odhner 1963)" baseAuthorityName="sensu Odhner" baseAuthorityYear="1963" class="Gastropoda" family="Tritoniidae" kingdom="Animalia" order="Nudibranchia" pageId="11" pageNumber="456" phylum="Mollusca" rank="family">
Tritoniidae (
<emphasis id="B97BEA88FF9B651CF970F8B6EB32F8C2" box="[1314,1365,1842,1865]" italics="true" pageId="11" pageNumber="456">sensu</emphasis>
<bibRefCitation id="EF9E4B6BFF9B651CF933F8B6ED06F8E2" author="Odhner NH" pageId="11" pageNumber="456" pagination="48 - 62" refId="ref26607" refString="Odhner NH. On the taxonomy of the family Tritoniidae (Mollusca: Opisthobranchia). The Veliger 1963; 6: 48 - 62." type="journal article" year="1963">Odhner 1963</bibRefCitation>
)
</taxonomicName>
has been recovered as paraphyletic in most phylogenetic studies with broad taxa sampling focusing on the family (
<bibRefCitation id="EF9E4B6BFF9B651CFF67F814ED90F82C" author="Huleu RE &amp; Mahguib J &amp; Gosliner TM" box="[821,1015,1935,1960]" pageId="11" pageNumber="456" pagination="215 - 222" refId="ref25435" refString="Huleu RE, Mahguib J, Gosliner TM, et al. Molecular evaluation of the phylogenetic position of the enigmatic species Trivettea papalotla (Bertsch) (Mollusca: Nudibranchia). Invertebrate Systematics 2015; 29: 215 - 222." type="journal article" year="2015">
Huleư
<emphasis id="B97BEA88FF9B651CFFD1F814EDD6F823" box="[899,945,1936,1960]" italics="true" pageId="11" pageNumber="456">et al</emphasis>
. 2015
</bibRefCitation>
,
<bibRefCitation id="EF9E4B6BFF9B651CF85AF814EB0DF823" author="Korshunova T &amp; Martynov A" box="[1032,1386,1936,1960]" pageId="11" pageNumber="456" pagination="0242103" refId="ref25588" refString="Korshunova T, Martynov A. Consolidated data on the phylogeny and evolution of the family Tritoniidae (Gastropoda: Nudibranchia) contribute to genera reassessment and clarify the taxonomic status of the neuroscience models Tritonia and Tochuina. PLoS One 2020; 15: e 0242103. hups: // doi. org / 10.1371 / journal. pone. 0242103" type="journal article" year="2020">Korshunova and Martynov 2020</bibRefCitation>
,
<bibRefCitation id="EF9E4B6BFF9B651CF929F814EDFDF84C" author="Rossi ME &amp; Avila C &amp; Moles J" pageId="11" pageNumber="456" pagination="559 - 573" refId="ref27227" refString="Rossi ME, Avila C, Moles J. Orange is the new white: taxonomic revision of Tritonia species (Gastropoda: Nudibranchia) from the Weddell Sea and Bouvet Island. Polar Biology 2021; 44: 559 - 573." type="journal article" year="2021">
Rossi
<emphasis id="B97BEA88FF9B651CFF78F834ED30F84C" box="[810,855,1967,1991]" italics="true" pageId="11" pageNumber="456">et al</emphasis>
. 2021
</bibRefCitation>
). Notably, the family was recovered with different topologies, depending on the composition of the superfamily
<taxonomicName id="4C0F4D19FF9C651BFCD3F8ACEF47F8CB" authorityName="Gray" authorityYear="1853" box="[129,288,1832,1856]" class="Gastropoda" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="457" phylum="Mollusca" rank="superFamily" superFamily="Proctonotoidea">Proctonotoidea</taxonomicName>
in our phylogenetic analyses (
<figureCitation id="13342A1FFF9C651BFE0FF8ACECF0F8CB" box="[605,663,1832,1856]" captionStart="Figure 1" captionStartId="9.[113,178,1772,1796]" captionTargetBox="[380,1196,144,1743]" captionTargetId="figure-8@9.[375,1197,144,1744]" captionTargetPageId="9" captionText="Figure 1. Bayesian phylogenetic trees based on the concatenated molecular data (COI + 16S + H3) of Tritoniidae (sensu Odhner 1963) with three distinct outgroup compositions: A, Proctonotoidea including Curnon granulosa (Curnonidae) and Dirona picta (Dironidae); B, Proctonotoidea including Antiopella barbarensis (Janolidae) and Antiopella capensis (Janolidae); C, no Proctonotoidea were included. Outgroup and clades are depicted in grey and Tritoniidae genera are independently coloured. PP are shown right/top and BS leħ/boưom. Unsupported branches are not labelled. Green circles indicate maximum support by both analyses." figureDoi="http://doi.org/10.5281/zenodo.8432391" httpUri="https://zenodo.org/record/8432391/files/figure.png" pageId="12" pageNumber="457">Fig. 1</figureCitation>
). A poorly supported relationship between
<taxonomicName id="4C0F4D19FF9C651BFD9AF8CCEC00F8EB" authorityName="Gray" authorityYear="1853" box="[456,615,1864,1888]" class="Gastropoda" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="467" phylum="Mollusca" rank="superFamily" superFamily="Proctonotoidea">Proctonotoidea</taxonomicName>
and
<taxonomicName id="4C0F4D19FF9C651BFEC5F8CCED6EF8EB" baseAuthorityName="sensu Odhner" baseAuthorityYear="1963" box="[663,777,1864,1888]" class="Gastropoda" family="Tritoniidae" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="457" phylum="Mollusca" rank="family">Tritoniidae</taxonomicName>
has previously only been suggested in a phylogeny based on transcriptomes (
<bibRefCitation id="EF9E4B6BFF9C651BFD63F802EC40F815" author="Goodheart JA &amp; Bazinet AL &amp; Collins AG" box="[305,551,1926,1950]" pageId="12" pageNumber="457" pagination="150196" refId="ref25067" refString="Goodheart JA, Bazinet AL, Collins AG, et al. Relationships within Cladobranchia (Gastropoda: Nudibranchia) based on RNA-Seq data: an initial investigation. Royal Society Open Science 2015; 2: 150196." type="journal article" year="2015">
Goodheart
<emphasis id="B97BEA88FF9C651BFDFDF803EF87F815" box="[431,480,1926,1950]" italics="true" pageId="12" pageNumber="457">et al</emphasis>
. 2015
</bibRefCitation>
). The only comprehensive phylogenetic study based on Sanger sequencing with members of the
<taxonomicName id="4C0F4D19FF9C651BFFBEF88DEAEAF8AA" authorityName="Gray" authorityYear="1853" box="[1004,1165,1801,1825]" class="Gastropoda" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="467" phylum="Mollusca" rank="superFamily" superFamily="Proctonotoidea">Proctonotoidea</taxonomicName>
,
<taxonomicName id="4C0F4D19FF9C651BF8C9F88DEB66F8AA" authorityName="Eliot" authorityYear="1907" box="[1179,1281,1801,1825]" class="Gastropoda" family="Tritoniidae" genus="Tritoniella" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="457" phylum="Mollusca" rank="genus">
<emphasis id="B97BEA88FF9C651BF8C9F88DEB66F8AA" box="[1179,1281,1801,1825]" italics="true" pageId="12" pageNumber="457">Tritoniella</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D19FF9C651BF969F88EEBF0F8AA" authorityName="Vayssiere" authorityYear="1877" box="[1339,1431,1802,1825]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="457" phylum="Mollusca" rank="genus">
<emphasis id="B97BEA88FF9C651BF969F88EEBF0F8AA" box="[1339,1431,1802,1825]" italics="true" pageId="12" pageNumber="457">Marionia</emphasis>
</taxonomicName>
, recovered
<taxonomicName id="4C0F4D19FF9C651BFFC2F8ACEA65F8CB" baseAuthorityName="sensu Odhner" baseAuthorityYear="1963" box="[912,1026,1832,1856]" class="Gastropoda" family="Tritoniidae" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="457" phylum="Mollusca" rank="family">Tritoniidae</taxonomicName>
as paraphyletic without a close relationship between
<taxonomicName id="4C0F4D19FF9C651BFFC8F8C3EA67F8D4" authorityName="Eliot" authorityYear="1907" box="[922,1024,1863,1887]" class="Gastropoda" family="Tritoniidae" genus="Tritoniella" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="457" phylum="Mollusca" rank="genus">
<emphasis id="B97BEA88FF9C651BFFC8F8C3EA67F8D4" box="[922,1024,1863,1887]" italics="true" pageId="12" pageNumber="457">Tritoniella</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D19FF9C651BF86BF8CCEAF2F8D4" authorityName="Vayssiere" authorityYear="1877" box="[1081,1173,1864,1887]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="457" phylum="Mollusca" rank="genus">
<emphasis id="B97BEA88FF9C651BF86BF8CCEAF2F8D4" box="[1081,1173,1864,1887]" italics="true" pageId="12" pageNumber="457">Marionia</emphasis>
</taxonomicName>
with other tritoniids (
<bibRefCitation id="EF9E4B6BFF9C651BF9D8F8C3EDC1F8F4" author="Rossi ME &amp; Avila C &amp; Moles J" pageId="12" pageNumber="457" pagination="559 - 573" refId="ref27227" refString="Rossi ME, Avila C, Moles J. Orange is the new white: taxonomic revision of Tritonia species (Gastropoda: Nudibranchia) from the Weddell Sea and Bouvet Island. Polar Biology 2021; 44: 559 - 573." type="journal article" year="2021">
Rossi
<emphasis id="B97BEA88FF9C651BFF6BF8ECED03F8F4" box="[825,868,1895,1919]" italics="true" pageId="12" pageNumber="457">et al</emphasis>
. 2021
</bibRefCitation>
). Phylogenetic studies that included
<taxonomicName id="4C0F4D19FF9C651BF97CF8E3EBF3F8F4" authorityName="Eliot" authorityYear="1907" box="[1326,1428,1895,1919]" class="Gastropoda" family="Tritoniidae" genus="Tritoniella" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="457" phylum="Mollusca" rank="genus">
<emphasis id="B97BEA88FF9C651BF97CF8E3EBF3F8F4" box="[1326,1428,1895,1919]" italics="true" pageId="12" pageNumber="457">Tritoniella</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D19FF9C651BFF6BF803EDF2F815" authorityName="Vayssiere" authorityYear="1877" box="[825,917,1927,1950]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="457" phylum="Mollusca" rank="genus">
<emphasis id="B97BEA88FF9C651BFF6BF803EDF2F815" box="[825,917,1927,1950]" italics="true" pageId="12" pageNumber="457">Marionia</emphasis>
</taxonomicName>
, but not members of the
<taxonomicName id="4C0F4D19FF9C651BF8FDF802EB37F815" authorityName="Gray" authorityYear="1853" box="[1199,1360,1926,1950]" class="Gastropoda" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="467" phylum="Mollusca" rank="superFamily" superFamily="Proctonotoidea">Proctonotoidea</taxonomicName>
, recovered a relationship between
<taxonomicName id="4C0F4D19FF9C651BF87EF821EAF5F836" authorityName="Eliot" authorityYear="1907" box="[1068,1170,1957,1981]" class="Gastropoda" family="Tritoniidae" genus="Tritoniella" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="457" phylum="Mollusca" rank="genus">
<emphasis id="B97BEA88FF9C651BF87EF821EAF5F836" box="[1068,1170,1957,1981]" italics="true" pageId="12" pageNumber="457">Tritoniella</emphasis>
</taxonomicName>
and other plateless tritoniids (
<bibRefCitation id="EF9E4B6BFF9D651AFC2EF880EF54F897" author="Valdes A &amp; Lundsten L &amp; Wilson NG" box="[124,307,1796,1820]" pageId="13" pageNumber="458" pagination="401 - 433" refId="ref27845" refString="Valdes A, Lundsten L, Wilson NG. Five new deep-sea species of nudibranchs (Gastropoda: Heterobranchia: Cladobranchia) from the Northeast Pacific. Zootaxa 2018; 4526: 401 - 433." type="journal article" year="2018">
Valdés
<emphasis id="B97BEA88FF9D651AFC95F881EE96F897" box="[199,241,1796,1820]" italics="true" pageId="13" pageNumber="458">et al</emphasis>
. 2018
</bibRefCitation>
,
<bibRefCitation id="EF9E4B6BFF9D651AFD12F880ECF2F897" author="Korshunova T &amp; Martynov A" box="[320,661,1796,1821]" pageId="13" pageNumber="458" pagination="0242103" refId="ref25588" refString="Korshunova T, Martynov A. Consolidated data on the phylogeny and evolution of the family Tritoniidae (Gastropoda: Nudibranchia) contribute to genera reassessment and clarify the taxonomic status of the neuroscience models Tritonia and Tochuina. PLoS One 2020; 15: e 0242103. hups: // doi. org / 10.1371 / journal. pone. 0242103" type="journal article" year="2020">Korshunova and Martynov 2020</bibRefCitation>
,
<bibRefCitation id="EF9E4B6BFF9D651AFEF0F880EEA3F8B7" author="Moles J &amp; Berning MI &amp; Hooker Y" pageId="13" pageNumber="458" pagination="107209" refId="ref26342" refString="Moles J, Berning MI, Hooker Y, et al. Due South: the evolutionary history of sub-Antarctic and Antarctic Tritoniidae nudibranchs. Molecular Phylogenetics and Evolution 2021; 162: 107209." type="journal article" year="2021">
Moles
<emphasis id="B97BEA88FF9D651AFEBBF881EEE4F8B7" italics="true" pageId="13" pageNumber="458">et al</emphasis>
. 2021
</bibRefCitation>
). This paưern was confirmed in our phylogenetic analyses: the presence of
<taxonomicName id="4C0F4D19FF9D651AFD06F8C7EF94F8D0" authorityName="Gray" authorityYear="1853" box="[340,499,1859,1883]" class="Gastropoda" kingdom="Animalia" order="Nudibranchia" pageId="13" pageNumber="467" phylum="Mollusca" rank="superFamily" superFamily="Proctonotoidea">Proctonotoidea</taxonomicName>
renders
<taxonomicName id="4C0F4D19FF9D651AFE1CF8C7ECA7F8D0" baseAuthorityName="sensu Odhner" baseAuthorityYear="1963" box="[590,704,1859,1883]" class="Gastropoda" family="Tritoniidae" kingdom="Animalia" order="Nudibranchia" pageId="13" pageNumber="458" phylum="Mollusca" rank="family">Tritoniidae</taxonomicName>
paraphyletic, with no relationships recovered between
<taxonomicName id="4C0F4D19FF9D651AFEDDF8E6EC92F8F1" authorityName="Eliot" authorityYear="1907" box="[655,757,1890,1914]" class="Gastropoda" family="Tritoniidae" genus="Tritoniella" kingdom="Animalia" order="Nudibranchia" pageId="13" pageNumber="458" phylum="Mollusca" rank="genus">
<emphasis id="B97BEA88FF9D651AFEDDF8E6EC92F8F1" box="[655,757,1890,1914]" italics="true" pageId="13" pageNumber="458">Tritoniella</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0F4D19FF9D651AFC23F806EEACF811" box="[113,203,1922,1946]" class="Gastropoda" family="Tritoniidae" genus="Tochuina" kingdom="Animalia" order="Nudibranchia" pageId="13" pageNumber="458" phylum="Mollusca" rank="genus">
<emphasis id="B97BEA88FF9D651AFC23F806EEACF811" box="[113,203,1922,1946]" italics="true" pageId="13" pageNumber="458">Tochuina</emphasis>
</taxonomicName>
and the marioniids with other members of the family (
<figureCitation id="13342A1FFF9D651AFC2EF825EE83F833" box="[124,228,1953,1977]" captionStart="Figure 1" captionStartId="9.[113,178,1772,1796]" captionTargetBox="[380,1196,144,1743]" captionTargetId="figure-8@9.[375,1197,144,1744]" captionTargetPageId="9" captionText="Figure 1. Bayesian phylogenetic trees based on the concatenated molecular data (COI + 16S + H3) of Tritoniidae (sensu Odhner 1963) with three distinct outgroup compositions: A, Proctonotoidea including Curnon granulosa (Curnonidae) and Dirona picta (Dironidae); B, Proctonotoidea including Antiopella barbarensis (Janolidae) and Antiopella capensis (Janolidae); C, no Proctonotoidea were included. Outgroup and clades are depicted in grey and Tritoniidae genera are independently coloured. PP are shown right/top and BS leħ/boưom. Unsupported branches are not labelled. Green circles indicate maximum support by both analyses." figureDoi="http://doi.org/10.5281/zenodo.8432391" httpUri="https://zenodo.org/record/8432391/files/figure.png" pageId="13" pageNumber="458">Fig. 1A, B</figureCitation>
). Moreover, this phenomenon is probably not due to any particular sequence included, as two distinct compositions of
<taxonomicName id="4C0F4D19FF9D651AFFD7F8A0EA43F8B7" authorityName="Gray" authorityYear="1853" box="[901,1060,1828,1852]" class="Gastropoda" kingdom="Animalia" order="Nudibranchia" pageId="13" pageNumber="467" phylum="Mollusca" rank="superFamily" superFamily="Proctonotoidea">Proctonotoidea</taxonomicName>
produced the same effect (
<figureCitation id="13342A1FFF9D651AF96FF8A0EBC5F8B0" box="[1341,1442,1828,1852]" captionStart="Figure 1" captionStartId="9.[113,178,1772,1796]" captionTargetBox="[380,1196,144,1743]" captionTargetId="figure-8@9.[375,1197,144,1744]" captionTargetPageId="9" captionText="Figure 1. Bayesian phylogenetic trees based on the concatenated molecular data (COI + 16S + H3) of Tritoniidae (sensu Odhner 1963) with three distinct outgroup compositions: A, Proctonotoidea including Curnon granulosa (Curnonidae) and Dirona picta (Dironidae); B, Proctonotoidea including Antiopella barbarensis (Janolidae) and Antiopella capensis (Janolidae); C, no Proctonotoidea were included. Outgroup and clades are depicted in grey and Tritoniidae genera are independently coloured. PP are shown right/top and BS leħ/boưom. Unsupported branches are not labelled. Green circles indicate maximum support by both analyses." figureDoi="http://doi.org/10.5281/zenodo.8432391" httpUri="https://zenodo.org/record/8432391/files/figure.png" pageId="13" pageNumber="458">Fig. 1A, B</figureCitation>
). Nevertheless, three-gene analyses are notoriously poor at providing robust family-level phylogenies because they suffer from problems with homology and variable region alignments (
<bibRefCitation id="EF9E4B6BFF9D651AFF67F825ED9FF832" author="Hallas JM &amp; Chichvarkhin A &amp; Gosliner TM" box="[821,1016,1953,1977]" pageId="13" pageNumber="458" pagination="171095" refId="ref25398" refString="Hallas JM, Chichvarkhin A, Gosliner TM. Aligning evidence: concerns regarding multiple sequence alignments in estimating the phylogeny of the Nudibranchia suborder Doridina. Royal Society Open Science 2017; 4: 171095." type="journal article" year="2017">
Hallas
<emphasis id="B97BEA88FF9D651AFFD0F826EDD5F832" box="[898,946,1953,1977]" italics="true" pageId="13" pageNumber="458">et al</emphasis>
. 2017
</bibRefCitation>
), and the possible relationship between
<taxonomicName id="4C0F4D19FF9E6519FCD3FA2FEF47FA48" authorityName="Gray" authorityYear="1853" box="[129,288,1451,1475]" class="Gastropoda" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="467" phylum="Mollusca" rank="superFamily" superFamily="Proctonotoidea">Proctonotoidea</taxonomicName>
and
<taxonomicName id="4C0F4D19FF9E6519FD32FA2FEFB5FA48" baseAuthorityName="sensu Odhner" baseAuthorityYear="1963" box="[352,466,1451,1475]" class="Gastropoda" family="Tritoniidae" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="459" phylum="Mollusca" rank="family">Tritoniidae</taxonomicName>
is not supported by recent transcriptome-based studies (
<bibRefCitation id="EF9E4B6BFF9E6519FDFDFA4FECEBFA68" author="Goodheart JA &amp; Bazinet AL &amp; Valdes A" box="[431,652,1483,1507]" pageId="14" pageNumber="459" pagination="1 - 14" refId="ref25106" refString="Goodheart JA, Bazinet AL, Valdes A, et al. Prey preference follows phylogeny: evolutionary dietary pauerns within the marine gastropod group Cladobranchia (Gastropoda: Heterobranchia: Nudibranchia). BMC Evolutionary Biology 2017; 17: 1 - 14." type="journal article" year="2017">
Goodheart
<emphasis id="B97BEA88FF9E6519FE77FA48EC2AFA68" box="[549,589,1483,1507]" italics="true" pageId="14" pageNumber="459">et al</emphasis>
. 2017
</bibRefCitation>
).
</paragraph>
<caption id="DF706612FF9C651BFCD3FA79EF57F952" ID-DOI="http://doi.org/10.5281/zenodo.8432399" ID-Zenodo-Dep="8432399" httpUri="https://zenodo.org/record/8432399/files/figure.png" pageId="12" pageNumber="457" startId="12.[129,194,1533,1557]" targetBox="[321,1281,144,1505]" targetPageId="12" targetType="figure">
<paragraph id="8BB0369AFF9C651BFCD3FA79EF57F952" blockId="12.[129,1472,1533,1753]" pageId="12" pageNumber="457">
<emphasis id="B97BEA88FF9C651BFCD3FA79EEBFF99E" bold="true" box="[129,216,1533,1557]" pageId="12" pageNumber="457">Figure 3.</emphasis>
Representative species of genera of
<taxonomicName id="4C0F4D19FF9C651BFE78FA79ECF3F99E" baseAuthorityName="sensu Odhner" baseAuthorityYear="1963" box="[554,660,1533,1557]" class="Gastropoda" family="Tritoniidae" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="457" phylum="Mollusca" rank="family">Tritoniidae</taxonomicName>
(with exception of
<taxonomicName id="4C0F4D19FF9C651BFF1EFA7AEDDDF99E" authorityName="Baba" authorityYear="1949" box="[844,954,1534,1557]" class="Gastropoda" family="Tritoniidae" genus="Paratritonia" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="457" phylum="Mollusca" rank="genus">
<emphasis id="B97BEA88FF9C651BFF1EFA7AEDDDF99E" box="[844,954,1534,1557]" italics="true" pageId="12" pageNumber="457">Paratritonia</emphasis>
</taxonomicName>
). A,
<taxonomicName id="4C0F4D19FF9C651BFFBBFA7AEAF5F99E" authorityName="Cuvier" authorityYear="1803" box="[1001,1170,1533,1557]" class="Gastropoda" family="Tritoniidae" genus="Tritonia" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="457" phylum="Mollusca" rank="species" species="hombergii">
<emphasis id="B97BEA88FF9C651BFFBBFA7AEAF5F99E" box="[1001,1170,1533,1557]" italics="true" pageId="12" pageNumber="457">Tritonia hombergii</emphasis>
</taxonomicName>
(MNCN15.05/93379) (Portaferry, Ireland); B,
<taxonomicName id="4C0F4D19FF9C651BFD36F99DEC67F9BA" baseAuthorityName="G. Johnston" baseAuthorityYear="1828" box="[356,512,1561,1585]" class="Gastropoda" family="Tritoniidae" genus="Candiella" higherTaxonomySource="GBIF" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="457" phylum="Mollusca" rank="species" species="plebeia">
<emphasis id="B97BEA88FF9C651BFD36F99DEC67F9BA" box="[356,512,1561,1585]" italics="true" pageId="12" pageNumber="457">Candiella plebeia</emphasis>
</taxonomicName>
(MNCN15.05/93393) (Portaferry, Ireland); C,
<taxonomicName id="4C0F4D19FF9C651BFF9FF99EEA1CF9BA" authorityName="Audouin" authorityYear="1826" box="[973,1147,1561,1585]" class="Gastropoda" family="Tritoniidae" genus="Tritoniopsis" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="457" phylum="Mollusca" rank="species" species="elegans">
<emphasis id="B97BEA88FF9C651BFF9FF99EEA1CF9BA" box="[973,1147,1561,1585]" italics="true" pageId="12" pageNumber="457">Tritoniopsis elegans</emphasis>
</taxonomicName>
(MB28-0045061) (Zavora, Mozambique); D,
<taxonomicName id="4C0F4D19FF9C651BFD7DF9B1EF8FF9C6" box="[303,488,1589,1613]" class="Gastropoda" family="Tritoniidae" genus="Tritonidoxa" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="457" phylum="Mollusca" rank="species" species="capensis">
<emphasis id="B97BEA88FF9C651BFD7DF9B1EF8FF9C6" box="[303,488,1589,1613]" italics="true" pageId="12" pageNumber="457">Tritonidoxa capensis</emphasis>
</taxonomicName>
(MNCN15.05/93399) (Scoưburg, South Africa); E,
<taxonomicName id="4C0F4D19FF9C651BFFB7F9B1EA0AF9C6" authorityName="Eliot" authorityYear="1907" box="[997,1133,1589,1613]" class="Gastropoda" family="Tritoniidae" genus="Tritoniella" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="457" phylum="Mollusca" rank="species" species="belli">
<emphasis id="B97BEA88FF9C651BFFB7F9B1EA0AF9C6" box="[997,1133,1589,1613]" italics="true" pageId="12" pageNumber="457">Tritoniella belli</emphasis>
</taxonomicName>
(Antarctica); F,
<taxonomicName id="4C0F4D19FF9C651BF95BF9B2EEC8F9E2" class="Gastropoda" family="Tritoniidae" genus="Marianina" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="457" phylum="Mollusca" rank="species" species="rosea">
<emphasis id="B97BEA88FF9C651BF95BF9B2EEC8F9E2" italics="true" pageId="12" pageNumber="457">Marianina rosea</emphasis>
</taxonomicName>
(French Polynesia); G,
<taxonomicName id="4C0F4D19FF9C651BFDC2F9D5EC3BF9E2" authorityName="Silva &amp; Pola &amp; Cervera" authorityYear="2023" baseAuthorityName="Bergh" baseAuthorityYear="1884" box="[400,604,1617,1641]" class="Gastropoda" family="Tritoniidae" genus="Myrella" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="457" phylum="Mollusca" rank="species" species="challengeriana" status="comb. nov.">
<emphasis id="B97BEA88FF9C651BFDC2F9D5EC3BF9E2" box="[400,604,1617,1641]" italics="true" pageId="12" pageNumber="457">Myrella challengeriana</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A24857F3FF9C651BFE33F9D5ECAEF9E2" box="[609,713,1617,1641]" pageId="12" pageNumber="457" rank="species">comb. nov.</taxonomicNameLabel>
(Bahía Camarones, Argentina); H,
<taxonomicName id="4C0F4D19FF9C651BF844F9D5EADDF9E2" authorityName="Bergh" authorityYear="1904" box="[1046,1210,1617,1641]" class="Gastropoda" family="Tritoniidae" genus="Tochuina" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="457" phylum="Mollusca" rank="species" species="gigantea">
<emphasis id="B97BEA88FF9C651BF844F9D5EADDF9E2" box="[1046,1210,1617,1641]" italics="true" pageId="12" pageNumber="457">Tochuina gigantea</emphasis>
</taxonomicName>
(Agnew Pass, USA); I,
<taxonomicName id="4C0F4D19FF9C651BFCD3F9E9EF57F90E" baseAuthorityName="Silva, Azevedo and Mauhews-Cascon" baseAuthorityYear="2014" box="[129,304,1645,1669]" class="Gastropoda" family="Tritoniidae" genus="Tritonicula" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="457" phylum="Mollusca" rank="species" species="khaleesi">
<emphasis id="B97BEA88FF9C651BFCD3F9E9EF57F90E" box="[129,304,1645,1669]" italics="true" pageId="12" pageNumber="457">Tritonicula khaleesi</emphasis>
</taxonomicName>
(MZUSP113165) (Fortaleza, Brazil); J,
<taxonomicName id="4C0F4D19FF9C651BFEE2F9EAED3CF90E" authorityName="Couthouy" authorityYear="1852" box="[688,859,1645,1669]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="457" phylum="Mollusca" rank="species" species="cucullata">
<emphasis id="B97BEA88FF9C651BFEE2F9EAED3CF90E" box="[688,859,1645,1669]" italics="true" pageId="12" pageNumber="457">Marionia cucullata</emphasis>
</taxonomicName>
(Golfo Nuevo, Argentina); (K)
<taxonomicName id="4C0F4D19FF9C651BF8D9F9EAEB4CF90E" authorityName="Eliot" authorityYear="1904" box="[1163,1323,1645,1669]" class="Gastropoda" family="Tritoniidae" genus="Marioniopsis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Nudibranchia" pageId="12" pageNumber="457" phylum="Mollusca" rank="species" species="levis">
<emphasis id="B97BEA88FF9C651BF8D9F9EAEB4CF90E" box="[1163,1323,1645,1669]" italics="true" pageId="12" pageNumber="457">Marioniopsis levis</emphasis>
</taxonomicName>
1 (MNHNC/ MB28-004895) (Nacala, Mozambique). Photo A and B, courtesy of B. Picton; photo C and K courtesy of Y. Tibiriça; photo D courtesy of V. Fraser; photo E courtesy of M. Caballer; photo F courtesy of G. Rouse; photo G and J, courtesy of N. Baưini; photo H courtesy of S. Thiebaud; photo I by F. Silva.
</paragraph>
</caption>
<caption id="DF706612FF9D651AFC23FA7CEF3BF95F" ID-DOI="http://doi.org/10.5281/zenodo.8432401" ID-Zenodo-Dep="8432401" httpUri="https://zenodo.org/record/8432401/files/figure.png" pageId="13" pageNumber="458" startId="13.[113,178,1528,1552]" targetBox="[309,1263,147,1498]" targetPageId="13" targetType="figure">
<paragraph id="8BB0369AFF9D651AFC23FA7CEBE8F933" blockId="13.[113,1430,1528,1749]" pageId="13" pageNumber="458">
<emphasis id="B97BEA88FF9D651AFC23FA7CEEAEF99B" bold="true" box="[113,201,1528,1552]" pageId="13" pageNumber="458">Figure 4.</emphasis>
Scanning electron micrographs of radulae of analysed key species of
<taxonomicName id="4C0F4D19FF9D651AFF1FFA7CEDD0F99B" authorityName="Lamarck" authorityYear="1809" box="[845,951,1528,1552]" pageId="13" pageNumber="462" rank="subFamily" subFamily="Tritoniinae">Tritoniinae</taxonomicName>
and
<taxonomicName id="4C0F4D19FF9D651AFFB6FA7CEA3FF99B" authority="Silva &amp; Pola &amp; Cervera, 2023" authorityName="Silva &amp; Pola &amp; Cervera" authorityYear="2023" box="[996,1112,1528,1552]" pageId="13" pageNumber="458" rank="subFamily" status="subfam. nov." subFamily="Marioniinae">Marioniinae</taxonomicName>
<taxonomicNameLabel id="A24857F3FF9D651AF80CFA7CEABEF99B" box="[1118,1241,1528,1552]" pageId="13" pageNumber="458" rank="subFamily">subfam. nov.</taxonomicNameLabel>
: A, tricuspid rachidian teeth and differentiated first lateral of
<taxonomicName id="4C0F4D19FF9D651AFE7DF991ECBFF9A7" authorityName="Cuvier" authorityYear="1803" box="[559,728,1556,1580]" class="Gastropoda" family="Tritoniidae" genus="Tritonia" kingdom="Animalia" order="Nudibranchia" pageId="13" pageNumber="458" phylum="Mollusca" rank="species" species="hombergii">
<emphasis id="B97BEA88FF9D651AFE7DF991ECBFF9A7" box="[559,728,1556,1580]" italics="true" pageId="13" pageNumber="458">Tritonia hombergii</emphasis>
</taxonomicName>
(ZMBN 83471); B, tricuspid rachidian teeth and differentiated first lateral with single accessory denticle (circled) of
<taxonomicName id="4C0F4D19FF9D651AFDAEF9B4ECF0F9C3" baseAuthorityName="G. Johnston" baseAuthorityYear="1828" box="[508,663,1584,1608]" class="Gastropoda" family="Tritoniidae" genus="Candiella" higherTaxonomySource="GBIF" kingdom="Animalia" order="Nudibranchia" pageId="13" pageNumber="458" phylum="Mollusca" rank="species" species="plebeia">
<emphasis id="B97BEA88FF9D651AFDAEF9B4ECF0F9C3" box="[508,663,1584,1608]" italics="true" pageId="13" pageNumber="458">Candiella plebeia</emphasis>
</taxonomicName>
(MNCN15.05/93393); C, unicuspid rachidian teeth and undifferentiated first lateral of
<taxonomicName id="4C0F4D19FF9D651AFC9AF9C9EF11F9EF" authorityName="Audouin" authorityYear="1826" box="[200,374,1612,1636]" class="Gastropoda" family="Tritoniidae" genus="Tritoniopsis" kingdom="Animalia" order="Nudibranchia" pageId="13" pageNumber="458" phylum="Mollusca" rank="species" species="elegans">
<emphasis id="B97BEA88FF9D651AFC9AF9C9EF11F9EF" box="[200,374,1612,1636]" italics="true" pageId="13" pageNumber="458">Tritoniopsis elegans</emphasis>
</taxonomicName>
(MNHNC/MB28-004506); D, tricuspid rachidian teeth and differentiated first lateral of
<taxonomicName id="4C0F4D19FF9D651AF891F9C8EB1AF9EF" box="[1219,1405,1612,1636]" class="Gastropoda" family="Tritoniidae" genus="Tritonidoxa" kingdom="Animalia" order="Nudibranchia" pageId="13" pageNumber="458" phylum="Mollusca" rank="species" species="capensis">
<emphasis id="B97BEA88FF9D651AF891F9C8EB1AF9EF" box="[1219,1405,1612,1636]" italics="true" pageId="13" pageNumber="458">Tritonidoxa capensis</emphasis>
</taxonomicName>
(MNCN15.05/93399); E, unicuspid rachidian teeth and undifferentiated first lateral of
<taxonomicName id="4C0F4D19FF9D651AFFF9F9ECEA28F90B" authorityName="Bergh" authorityYear="1904" box="[939,1103,1640,1664]" class="Gastropoda" family="Tritoniidae" genus="Tochuina" kingdom="Animalia" order="Nudibranchia" pageId="13" pageNumber="458" phylum="Mollusca" rank="species" species="gigantea">
<emphasis id="B97BEA88FF9D651AFFF9F9ECEA28F90B" box="[939,1103,1640,1664]" italics="true" pageId="13" pageNumber="458">Tochuina gigantea</emphasis>
</taxonomicName>
(NHMD-115967); F, tricuspid rachidian teeth and differentiated first lateral with several accessory denticles (circled) of
<taxonomicName id="4C0F4D19FF9D651AFFE6F900EAF5F917" baseAuthorityName="Gosliner and Ghiselin" baseAuthorityYear="1987" box="[948,1170,1668,1692]" class="Gastropoda" family="Tritoniidae" genus="Tritonicula" kingdom="Animalia" order="Nudibranchia" pageId="13" pageNumber="458" phylum="Mollusca" rank="species" species="hamnerorum">
<emphasis id="B97BEA88FF9D651AFFE6F900EAF5F917" box="[948,1170,1668,1692]" italics="true" pageId="13" pageNumber="458">Tritonicula hamnerorum</emphasis>
</taxonomicName>
(MNCN15.05/93378); G, tricuspid rachidian teeth and differentiated first lateral of
<taxonomicName id="4C0F4D19FF9D651AFEDAF925ED27F933" authorityName="Risso" authorityYear="1818" box="[648,832,1696,1720]" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="13" pageNumber="458" phylum="Mollusca" rank="species" species="blainvillea">
<emphasis id="B97BEA88FF9D651AFEDAF925ED27F933" box="[648,832,1696,1720]" italics="true" pageId="13" pageNumber="458">Marionia blainvillea</emphasis>
</taxonomicName>
(NHMD115968); Scale bars: A, D, E = 60 μm; B, C = 22 μm;
</paragraph>
<paragraph id="8BB0369AFF9D651AFC23F939EF3BF95F" blockId="13.[113,1430,1528,1749]" box="[113,348,1725,1749]" pageId="13" pageNumber="458">F = 20 μm; G = 110 μm.</paragraph>
</caption>
<caption id="DF706612FF9E6519FCD3FB6AEC11FAFD" ID-DOI="http://doi.org/10.5281/zenodo.8432405" ID-Zenodo-Dep="8432405" httpUri="https://zenodo.org/record/8432405/files/figure.png" pageId="14" pageNumber="459" startId="14.[129,194,1262,1286]" targetBox="[164,1437,147,1230]" targetPageId="14" targetType="figure">
<paragraph id="8BB0369AFF9E6519FCD3FB6AEC11FAFD" blockId="14.[129,1436,1262,1398]" pageId="14" pageNumber="459">
<emphasis id="B97BEA88FF9E6519FCD3FB6AEEBFFA8D" bold="true" box="[129,216,1262,1286]" pageId="14" pageNumber="459">Figure 5.</emphasis>
Digestive system arrangements of analysed key species of
<taxonomicName id="4C0F4D19FF9E6519FEABFB6AED04FA8D" authorityName="Lamarck" authorityYear="1809" box="[761,867,1262,1286]" pageId="14" pageNumber="462" rank="subFamily" subFamily="Tritoniinae">Tritoniinae</taxonomicName>
and
<taxonomicName id="4C0F4D19FF9E6519FFC2FB6AEA63FA8D" authority="Silva &amp; Pola &amp; Cervera, 2023" authorityName="Silva &amp; Pola &amp; Cervera" authorityYear="2023" box="[912,1028,1262,1286]" pageId="14" pageNumber="459" rank="subFamily" status="subfam. nov." subFamily="Marioniinae">Marioniinae</taxonomicName>
<taxonomicNameLabel id="A24857F3FF9E6519F858FB6AEAE5FA8D" box="[1034,1154,1262,1286]" pageId="14" pageNumber="459" rank="subFamily">subfam. nov.</taxonomicNameLabel>
, dorsal view: A,
<taxonomicName id="4C0F4D19FF9E6519F949FB6BEEBCFAA9" authorityName="Cuvier" authorityYear="1803" class="Gastropoda" family="Tritoniidae" genus="Tritonia" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="459" phylum="Mollusca" rank="species" species="hombergii">
<emphasis id="B97BEA88FF9E6519F949FB6BEEBCFAA9" italics="true" pageId="14" pageNumber="459">Tritonia hombergii</emphasis>
</taxonomicName>
(ZMBN 83471); B,
<taxonomicName id="4C0F4D19FF9E6519FDCCFA8EEC5DFAA9" baseAuthorityName="G. Johnston" baseAuthorityYear="1828" box="[414,570,1290,1314]" class="Gastropoda" family="Tritoniidae" genus="Candiella" higherTaxonomySource="GBIF" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="459" phylum="Mollusca" rank="species" species="plebeia">
<emphasis id="B97BEA88FF9E6519FDCCFA8EEC5DFAA9" box="[414,570,1290,1314]" italics="true" pageId="14" pageNumber="459">Candiella plebeia</emphasis>
</taxonomicName>
(MNCN15.05/93393); C,
<taxonomicName id="4C0F4D19FF9E6519FF10FA8FED84FAA9" authorityName="Er. Marcus and Ev. Marcus" authorityYear="1970" box="[834,995,1290,1314]" class="Gastropoda" family="Tritoniidae" genus="Tritoniopsis" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="459" phylum="Mollusca" rank="species" species="flydis">
<emphasis id="B97BEA88FF9E6519FF10FA8FED84FAA9" box="[834,995,1290,1314]" italics="true" pageId="14" pageNumber="459">Tritoniopsis flydis</emphasis>
</taxonomicName>
(GR7-DMC00040); D,
<taxonomicName id="4C0F4D19FF9E6519F898FA8EEEADFAB5" class="Gastropoda" family="Tritoniidae" genus="Tritonidoxa" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="459" phylum="Mollusca" rank="species" species="capensis">
<emphasis id="B97BEA88FF9E6519F898FA8EEEADFAB5" italics="true" pageId="14" pageNumber="459">Tritonidoxa capensis</emphasis>
</taxonomicName>
(MNCN15.05/93399); E,
<taxonomicName id="4C0F4D19FF9E6519FD98FAA2EC09FAB5" authorityName="Bergh" authorityYear="1904" box="[458,622,1318,1342]" class="Gastropoda" family="Tritoniidae" genus="Tochuina" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="459" phylum="Mollusca" rank="species" species="gigantea">
<emphasis id="B97BEA88FF9E6519FD98FAA2EC09FAB5" box="[458,622,1318,1342]" italics="true" pageId="14" pageNumber="459">Tochuina gigantea</emphasis>
</taxonomicName>
(NHMD-115967); F,
<taxonomicName id="4C0F4D19FF9E6519FF17FAA2EA44FAB5" baseAuthorityName="Gosliner and Ghiselin" baseAuthorityYear="1987" box="[837,1059,1318,1342]" class="Gastropoda" family="Tritoniidae" genus="Tritonicula" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="459" phylum="Mollusca" rank="species" species="hamnerorum">
<emphasis id="B97BEA88FF9E6519FF17FAA2EA44FAB5" box="[837,1059,1318,1342]" italics="true" pageId="14" pageNumber="459">Tritonicula hamnerorum</emphasis>
</taxonomicName>
(MNCN15.05/93378); G,
<taxonomicName id="4C0F4D19FF9E6519F97FFAA3EEB9FAD1" authorityName="Risso" authorityYear="1818" class="Gastropoda" family="Tritoniidae" genus="Marionia" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="459" phylum="Mollusca" rank="species" species="blainvillea">
<emphasis id="B97BEA88FF9E6519F97FFAA3EEB9FAD1" italics="true" pageId="14" pageNumber="459">Marionia blainvillea</emphasis>
</taxonomicName>
(NHMD115968). Abbreviations: a, anus; aodg, anterior ODG complex; e, esophagus; i, intestine; odg, ovotestis-digestive gland complex; s, stomach; podg, posterior ODG complex.
</paragraph>
</caption>
<paragraph id="8BB0369AFF9E6519FCCEFA6EEB6AF9EB" blockId="14.[129,778,1451,1977]" lastBlockId="14.[825,1474,1451,1632]" pageId="14" pageNumber="459">
Based on the chosen phylogenetic hypothesis (
<figureCitation id="13342A1FFF9E6519FEC1FA6EECA8F989" box="[659,719,1514,1538]" captionStart="Figure 2" captionStartId="10.[129,194,1719,1743]" captionTargetBox="[325,1278,148,1689]" captionTargetId="figure-6@10.[321,1281,144,1692]" captionTargetPageId="10" captionText="Figure 2. Bayesian phylogenetic tree based on the concatenated molecular data (COI + 16S + H3) of Tritoniidae (sensu Odhner 1963). Outgroup taxa are depicted in grey.Tritoniidae genera are independently coloured, and branches with several replicates for each species hypothesis were collapsed for clarity.Names in bold are species which sequences were generated for this study with the number of sequenced specimens showed in brackets. PP are shown right/top and BS leħ/boưom.Unsupported branches are not labelled. Green circles indicate maximum support by both analyses. Solid vertical bars indicate specimens grouped as species entities based on COI dataset. Empty vertical bars indicate the specimens grouped as species based on the 16S dataset." figureDoi="http://doi.org/10.5281/zenodo.8432395" httpUri="https://zenodo.org/record/8432395/files/figure.png" pageId="14" pageNumber="459">Fig. 2</figureCitation>
) and morphological analysis, the traditional family
<taxonomicName id="4C0F4D19FF9E6519FE3AF98DECBDF9AA" baseAuthorityName="sensu Odhner" baseAuthorityYear="1963" box="[616,730,1545,1569]" class="Gastropoda" family="Tritoniidae" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="459" phylum="Mollusca" rank="family">Tritoniidae</taxonomicName>
is a non-monophyletic group primarily divided into plateless (clade A + B) and plate-bearing species (clade C). Thus, we propose the division of tritoniids into two new monophyletic subfamilies,
<taxonomicName id="4C0F4D19FF9E6519FCD3F902EE9AF915" authorityName="Lamarck" authorityYear="1809" box="[129,253,1670,1694]" pageId="14" pageNumber="462" rank="subFamily" subFamily="Tritoniinae">
<emphasis id="B97BEA88FF9E6519FCD3F902EE9AF915" bold="true" box="[129,253,1670,1694]" pageId="14" pageNumber="459">Tritoniinae</emphasis>
</taxonomicName>
and
<emphasis id="B97BEA88FF9E6519FD6AF902EC3EF914" bold="true" box="[312,601,1670,1695]" pageId="14" pageNumber="459">
<taxonomicName id="4C0F4D19FF9E6519FD6AF902EFD8F915" authority="Silva &amp; Pola &amp; Cervera, 2023" authorityName="Silva &amp; Pola &amp; Cervera" authorityYear="2023" box="[312,447,1670,1694]" pageId="14" pageNumber="459" rank="subFamily" status="subfam. nov." subFamily="Marioniinae">Marioniinae</taxonomicName>
<taxonomicNameLabel id="A24857F3FF9E6519FD98F902EC3EF914" box="[458,601,1670,1695]" pageId="14" pageNumber="459" rank="subFamily">subfam. nov.</taxonomicNameLabel>
</emphasis>
, to reflect these new findings, while preserving the rank of
<taxonomicName id="4C0F4D19FF9E6519FE19F922ECDAF935" baseAuthorityName="sensu Odhner" baseAuthorityYear="1963" box="[587,701,1702,1726]" class="Gastropoda" family="Tritoniidae" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="459" phylum="Mollusca" rank="family">Tritoniidae</taxonomicName>
within
<taxonomicName id="4C0F4D19FF9E6519FCD3F941EF7FF956" authorityName="Willan and Morton" authorityYear="1984" box="[129,280,1733,1757]" class="Gastropoda" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="456" phylum="Mollusca" rank="subOrder" subOrder="Cladobranchia">Cladobranchia</taxonomicName>
based on two well-established synapomorphies: (1) a retractable and sheathed palmate rhinophore consisting of a smooth club surrounded by a distal circle of vertically branched processes, and (2) one row of dorsolateral, arborescent secondary gills (Ev.
<bibRefCitation id="EF9E4B6BFF9E6519FD2BF8C7EF9AF8D0" author="Marcus Ev" box="[377,509,1859,1883]" pageId="14" pageNumber="459" pagination="177 - 214" refId="ref25994" refString="Marcus Ev. The Western Atlantic Tritoniidae. Boletim de Zoologia, Universidade de Sao Paulo 1983; 6: 177 - 214." type="journal article" year="1983">Marcus 1983</bibRefCitation>
). This reassessment is also consistent with the current conflicting molecular evidence for the monophyly of
<taxonomicName id="4C0F4D19FF9E6519FD12F805EFD5F812" baseAuthorityName="sensu Odhner" baseAuthorityYear="1963" box="[320,434,1921,1945]" class="Gastropoda" family="Tritoniidae" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="459" phylum="Mollusca" rank="family">Tritoniidae</taxonomicName>
and its sister-group, which could be
<taxonomicName id="4C0F4D19FF9E6519FCCDF825EF69F832" authorityName=", Aeolidioidea Gray" authorityYear="1827" box="[159,270,1953,1977]" class="Gastropoda" family="Arminidae" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="459" phylum="Mollusca" rank="family">Arminidae</taxonomicName>
,
<taxonomicName id="4C0F4D19FF9E6519FD4AF825EFD2F832" authorityName="Allman" authorityYear="1845" box="[280,437,1953,1977]" class="Gastropoda" family="Dendronotidae" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="459" phylum="Mollusca" rank="family">Dendronotidae</taxonomicName>
or some proctonotoidean family. Henceforth, the term tritoniids will be used in this study to refer to all species of
<taxonomicName id="4C0F4D19FF9E6519FF8AFA4FEA2BFA68" baseAuthorityName="sensu Odhner" baseAuthorityYear="1963" box="[984,1100,1483,1507]" class="Gastropoda" family="Tritoniidae" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="459" phylum="Mollusca" rank="family">Tritoniidae</taxonomicName>
, whilst plate less tritoniids will refer only to species recovered in the
<taxonomicName id="4C0F4D19FF9E6519F8DCFA6EEB67F989" authorityName="Lamarck" authorityYear="1809" box="[1166,1280,1514,1538]" pageId="14" pageNumber="462" rank="subFamily" subFamily="Tritoniinae">Tritoniinae</taxonomicName>
and plate-bearing tritoniids or marioniids will refer only to species recovered in the
<taxonomicName id="4C0F4D19FF9E6519FF31F9ADED84F9CA" authorityName="Silva &amp; Pola &amp; Cervera" authorityYear="2023" box="[867,995,1577,1601]" pageId="14" pageNumber="467" rank="subFamily" subFamily="Marioniinae">Marioniinae</taxonomicName>
. It is not possible to assign
<taxonomicName id="4C0F4D19FF9E6519F95BF9ADEB1CF9CA" baseAuthorityName="sensu Odhner" baseAuthorityYear="1963" box="[1289,1403,1577,1601]" class="Gastropoda" family="Tritoniidae" kingdom="Animalia" order="Nudibranchia" pageId="14" pageNumber="459" phylum="Mollusca" rank="family">Tritoniidae</taxonomicName>
to the superfamily
<taxonomicName id="4C0F4D19FF9E6519FFE5F9CCEA50F9EB" authorityName="Lamarck" authorityYear="1809" box="[951,1079,1608,1632]" pageId="14" pageNumber="459" rank="superFamily" superFamily="Tritonioidea">Tritonioidea</taxonomicName>
based on our results.
</paragraph>
</subSubSection>
</treatment>
</document>